IPOMOPSIS RAMOSA (POLEMONIACEAE), A NEW SPECIES FROM SOUTHWESTERN COLORADO Al Schneider V 19049 Road Lewis, Colorado 81327 John Bregar 506 Hillcrest Drive, Durango, Colorado 81301 ABSTRACT & Ipomopsis ramosa Schneider Bregar, sp. nov., is described from red sandstones, siltstones, and shales of the Permian Cutler Formation in two side canyons of the extensive Dolores River Canyon in southwest Colorado. In each side canyon, Ipomopsis ramosa represented by many is thousands of plants, where they grow on steep slopes in sunny meadows and open forests in moist but well-drained habitat between 8200 and 9200 feet elevation, with Picea pungens, Pseudotsuga menziesii, and Populus tremuloides. Ipomopsis ramosa most closely resembles sancti-spiritus and /. Ipomopsis tenuituba subsp, tenuituba, but ramosa significantly different from both in /, is morphology and geography and justifiably considered a distinct species. is KEY WORDS: Ipomopsis ramosa, Ipomopsis sancti-spiritus, Ipomopsis tenuituba, Polemoniaceae, Colorado endemics Biological exploration of San Juan National Forest in Montezuma County, Colorado, by Al and Betty Schneider in 2006 has brought to light a previously unknown species of Ipomopsis, described here as Ipomopsis ramosa. & Ipomopsis ramosa Schneider Bregar, sp. nov. Figs. 1-7. TYPE: USA. Colorado. Montezuma NE Co.: San Juan National Forest, ca. 28 miles of the town of Dolores, T38N, R11W, Sections 4, 5, 9, and 10; from 37° 35.84-108° 6.54' to 37° 34.70-108° 3.49'; exposure of 35% red sandstones, siltstones, and shales of the Permian Cutler formation, ca. slope, with a SW, S, or SE orientation at 8200-9200 feet (2500-2800 meters); plants with open, erect, multiple stems and corollas heavily streaked and lightly dotted with coral pink; 29 Jun 2006, A. Schneider 5085 with Betty Schneider (holotype: COLO isotypes: SBBG, RSA, MO, NY, RM UNM, US, ASU, BRY, FLD). Several other unnumbered collections were sent to experts for study. Otherwise no further collections ofIpomopsis ramosa have been made. Ipomopsi sancti-spiritus similis sed differt caulibus brevioribus maxinie ramosis, corollis Ipomopsi glandulosis albis corallini-striatis, aristis calycis brevioribus, et stylis longioribus. tenuitubae subsp. tenuitubae similis sed differt habitu breviore plus ramoso et corollis ac stylis brevioribus. cm Biennials to perennials from stout taproots to 24 long, with stout, sometimes branched mm caudices to 6 cm, up to 3 thick Stems: elect, commonly multi-stemmed and usually multi- mm 3— branched, main stems 25-44 cm tall, \ thick, stiff, often red, branched 4-9(-13) times from mm base and along main stem, rarely simple, branches 1-2 thick, often to the height of the main stem(s), moderately villous and glandular proximally, moderately villous and increasingly glandular distally, becoming moderately villous and highly glandular. Leaves: basal rosette prominent, 5-9 cm Bregar: Ipomopsis n d cm in diameter, leaves 2.5- 4.5 long, very sparsely villous, especially along rachis, leaf segments cm mucronate, withered black at anthesis; cauline leaves pmnatisect with linear lobes 2.5 long, proximal 3-5 cm long, reduced gradually distally; center of adaxial surfaces and center of lobes often strongly to lightly villous, margins commonly lightly glandular, sometimes also villous, both surfaces sparsely glandular, becoming strongly glandular distally. The glandularity produces a very noticeable and pleasant spicy fragrance. Inflorescence: occupying distal Vi to Va of the plant, of numerous pedicelled, open-paniculate clusters, the basic unit composed of 4-7 flowers, tending to be secund; mm mm pedicels at anthesis commonly 2-5 long, peduncles 20-30 long; each flower subtended by a 3^1 mm, single entire, linear, strong!) glandular biaet: bia^K folraccou'*. the uppermost bracts entire cm and linear, lower increasingly pinnatifid with linear lobes often entire, less than 2.5 long, often mm strongly glandular and lightly villous. Calyx: densely glandular, 6-7 long, tube ribs green and7 or mm purple, membrane connecting ribs white or purple; lobes equal, green, 2-4(-6) long, linear- lanceolate, attenuate with a non-herbaceous spinulose tip. Corolla: salverform, slightly translucent mm allowing colors to show through; tube straight or occasionally slightly curved, 14-17 long mm mm [measured from base of tube to sinuses of lobes], 0.7-1 in diameter at base, 1.7-2 in diameter at orifice, moderately to heavily glandular-puberulent, heavily coral pink streaked on white background, infrequently completely coral pink, fading abruptly to all white within the caryx; lobes 8 mm long, attenuate, spreading or slightly to moderately reflexed, the adaxial surface of the lobes white with scattered coral pink dots, becoming light coral pink and coral pink dotted toward the apex; abaxial surface white with numerous wide coral pink streaks. In bud, the dorsal surface of the lobes From appears orange, in contrast to the coral pink tube. a distance the overall appearance of the corolla salmon though close-up examination shows to be white with coral pink streaks and dots. is it Stamens: deeply included, unequal in length, unequally inserted on the tube; free part of filaments mm mm mm 0.1-1 long; anther sac 0.7 long by 0.5 wide, basifixed, vertical in tube; anther sac on mm 7- lowest stamen in tube 5 to 7 above base of the tube; anther sac on highest stamen in tube is is mm mm 10 above base of the tube; pollen yellow. Carpels: 2-3.5 long (excluding the stigmas), style mm very deeply included, stigma lobes 0.5 long, usually terminating below the anthers at anthesis. mm mm mm Fruit: obpyriform capsules, 3^1 long x 1.2 wide and green when corolla abscises; 5-6 mm long x 1.5 wide and tan at maturity. Seeds: 2-3 per locule, cylindric, young seeds bright green, 1 mm mm mm long x 0.3 wide, encased in milky, gelatinous substance; mature seeds dull green, 2+ mm now long x 0.5-1 wide, encased in the buff-colored, solidified gelatinous substance; seed coat mucilaginous when wet. Chromosome number unknown. The format of the description generally matches that of Wilken and Fletcher (1988) and Wilken and Porter (2005). The specific epithet, "ramosa," means "many-branched" and was chosen because the We branching of this species is a prominent and distinguishing visual characteristic. suggest the common name "Coral Ipomopsis." new Within several hundred miles of the populations of the species are four Ipomopsis species that superficially resemble but that differ from in significant morphological, geographical, and it it ecological characteristics. common Ipomopsis aggregate! (Pursh) Grant, very in the immediate vicinity of ramosa, /. is much markedly different in corolla color (usually scarlet) and in larger dimensions: its its it mm mm mm grows to 100 cm tall with corolla tubes to 30 long, styles 6-18 long, calyces 3-6 mm long, and capsules 8-12 long. Some details from Wilken and Porter (2005). Ipomopsis arizonica (Greene) Wherry resembles aggregata in a number of morphological /. its features and has been considered a variety of that species by Cronquist et al. (1984) and Welsh et al. (2008). It is found 250^150 miles southwest of the Colorado canyons in which /. ramosa is . Bregar: Ipomopsis n d found and is significantly distinct from /. ramosa having mostly single stems, glabrous to mm sparingly glandular upper internode pubescence, calyx lobes 1-3 long, scarlet corolla, stamens attached at the same level, style length 4.5-10 mm, stigma at same level as anthers, and habitat of pinyon-juniper or scrub-oak woodland. Most details from Wilken and Fletcher Ipomopsis tenuituba (Rydb.) V.E. Grant has its closest populations in the Abajo Mountains, Utah, 70 miles west of the /. ramosa populations. Its corollas are commonly satin white, salmon, lavender, or pink and it is significantly larger man /. ramosa in many of its dimensions. mm mm It grows to 100 cm tall with corolla tubes 25-45 long, throat 2-\ wide, anthers included mm to slightly exserted, stigma slightly exsertedto included, styles are 16-35 long, and capsules mm 5-8 long. 1. tenuituba grows on a variety of substrates in high montane meadows and open aspen and conifer forests. Some details from Wilken and Porter (2005), Wilken (1993), and Grant and Wilken 1988. (Note: a Dolores Co., Colorado, specimen identified by Moore in 1995 as tenuituba (RM 714481) was, according to Ron Hartman, misidentified. The specimen I. is /. & Ipomopsis sancti-spiritus Wilken Fletcher found 180 miles southeast of ramosa in one is /. New small, rare, and endangered population in San Miguel Co., Mexico. It grows from 30-80 cm tall and has solitary stems. Corollas are uniformly pink to hot pink, glabrous; tubes are 15- mm mm mm 18 long; calyces are green, white, and purple, 5-7 long; styles are 1.3-2 long; mm stamens are deeply included; capsules are 4-5 long. It is found from 7600 feet to 8200 feet on substrate derived from the Terrero and Espiritu Santo Formation, primarily a sandy to pebbly NMRPTC limestone conglomerate. Most details from Wilken and Fletcher (1988), (2011), and personal communications from Phil Tonne and Bob Sivinski. Ipomopsis ramosa grows from 25—44 cm and commonly has multiple stems almost tall always with numerous branches. Corollas are white with short streaks and dots of coral pink; tubes mm are slightly translucent, strongly glandular, 14-17 long; calyces are green, white, and purple, 6-7 mm mm mm long; styles are 2.5-4 long; stamens are deeply included; capsules are 5-6 long. It occurs from 8200-9200 feet on red sandstones, siltstones, and shales of the Permian Cutler Formation. The species most similar to Ipomopsis ramosa are sancti-spiritus and tenuituba subsp. I. /. tenuituba. The three are similar in their generally dense glandularity, open to narrow, often 1-sided inflorescences, and short-pedicelled to subsessile flowers in lateral pedunculate clusters. ramosa /. and sancti-spiritus are further similar in their short, pink or coral pink and white corollas, very short /. styles, calyx, dimensions, and deeply included anthers. Ipomopsis ramosa and tenuituba subsp. /. tenuituba are imlh^i bmulai in then dtn.se upper internode glandularity, calyx colors, dotted pattern on corolla lobes, glandularity of corolla, and shape of corolla lobes. Differences between Ipomopsis ramosa and sancti-spiritus and tenuituba ssp. tenuituba /. I. are summarized in the following pair of couplets and in Table 1 Stems commonly multiple, frequently branched, 25^-14 cm densely glandular and moderately 1. tall, villous upward; corolla tubes slightly translucent, heavily coral pink streaked, ventral surface of lobes lightly coral pink dotted, dorsal surface moderately coral pink streaked, moderately to heavily mm mm glandular, tubes 14-17 mm; style 2.5-4 at anthesis; stamens deeply inserted 5-10 above the mm base of the tube; filaments 0. 1-1 long, straight; habitat over sandstones, siltstones, and shales at 8200-9200 Ipomopsis ramosa feet Bregar: Ipomopsis n d 30-80 cm Stems mostly simple, occasionally branched, denser} gbndular upward; corollas 1. tall, mm uniformly pink to hot pink, glabrous, tubes 15-18 mm; style 1.3-2 at anthesis; stamens deeply habitat over limestone at y; Ipomopsis sancti-spiritus 1. Stems commonly multiple, frequently branched, 25-44 cm tall, not woody at base, densely mm glandular and moderately villous upward; calyx awn 0.2-0.3 long; corolla tubes slightly translucent, heavily coral pink streaked, ventral surface of lobes lightly coral pink dotted, dorsal surface moderately coral pink streaked, moderately to heavily glandular, tubes 14-17 mm, throat 1.7- mm mm mm 2 wide; style 2.5-4 at anthesis, very deeply inserted; stamens deeply inserted 5-10 mm above the base of the tube; filaments 0. 1-1 long, straight; habitat over sandstones, siltstones, and 8200-9200 Ipomopsis ramosa shales at feet 1. Stems usually simple, occasionally branched, 35-100 cm tall, often woody at base, densely mm glandular upward; calyx awn 0.6-0.9 long; corolla tubes and lobes satin, commonly white, salmon, lavender, or pink, lightly to moderately glandular, ventral surface of lobes occasionally mm mm dotted, tubes 25^45 mm, throat 2-4 wide; style 16-35 at anthesis. stigma may be slightly mm exserted; stamens inserted to 6 or some or all exserted to 2 mm; filaments 1-1.5, often kinked; habitat various montane meadows and open aspen and conifer forests Ipomopsis tenuitubasubsp. tenuituba Habitat and occurrence. Ipomopsis ramosa currently known exclusively from two large populations, one in type is its locale, a five mile stretch of the Roaring Fork Canyon, 28 miles northeast of Dolores, and the other in — a one mile stretch of Wildcat Canyon, thee miles north of the Roaring Fork both in Montezuma Co., Colorado. The new species represented by thousands of plants in each canyon and among is it is the more abundant species in its habitat. Explorations in nearby canyons with similar Cutler Formation and similar southerly orientations have so far revealed no further populations; additional searches will be needed to determine if there are more populations of the new species on the high, steep slopes in Roaring Fork and Wildcat Canyons and if there are populations in nearby canyons. Ipomopsis ramosa occurs most often as well separated individual plants but does it occasionally occur in small clusters. Plants most often occur on sparsely vegetated soils on steep slopes (ca. 35%) with a southwest, south, or southeast orientation. The new species grows in well drained soils and on rocky ledges derived from red sandstones, siltstones, and shales of the Permian Cutler Formation. 'The plants flower from early June into September. Over the extent of habitat, L its ramosa very uniform morphology. in is At the type locality Ipomopsis ramosa grows with the following dominant woody plants: Picea pungens, Pseudotsuga menziesii, Populus tremuloides, Juniperus communis, and Rosa woodsii. Less common woody plants; Pinus flexilis, Quercus gambelii, Amelanchier alnifolia, Lonicera involucrata, Populus angustifolia, and Salix spp. Associated dominant forbs: Heterotheca villosa, Ipomopsis aggregata, Packera neomexicana, Fragaria virginiana, Lathyrus lanszwertii var. leucanthus, Geranium caespitosum, Geranium richardsonii, Cymopterus lemmonii, Penstemon sirictus. Oenothera elata, Galium boreale, Achillea millefolium, Lithospermum multiflorum, Erigeron Apocynum speciosus, Erigeron flagellaris, Potentilla pulcherrima, androsaemifolium, Castilleja miniata, Oreochrysum parryi, Hedysarum occidentale, Eriogonum ovalifolium, Valeriana edulis, Viola adunca, and various grasses. : Bregar: Ipomopsis n d Table 1. Ipomopsis r compared to Ipomopsis sancti-spiritus and Ipomopsis tenuituba ssp. i The most tenuituba. cant contrasts are in bold. ramosa sanet.-sp.ntus */. tenuituba ssp. /. /. tenuituba New Location. sw Colorado; spruce, central Mexico; eastern California through habitat, douglas aspen; steep ponderosa, aspen: steep, Nevada and Utah to fir, SW elevation SE, S, or slopes; SSW-facing slopes; Arizona and central 8200-9200 on red 7600-8200 on Colorado; meadows and feet; feet; sandstones, siltstones, substrate derived from open woods in the high and shales of the Permian the Terrero and Espiritu montane aspen and conifer Cutler Formation Santo Formation, zone on various substrates consisting primarily of a sandy pebbly to limestone conglomerate Variability highly uniform relatively uniform highly variable in color of corolla (even within a single color and population) and morphology dimensions Height of 25-^4 cm 30-80 cm 35-100 cm plant Branching commonly branched occasionally branched simple to branched of plant from base and' or main from base stem mm mm mm 20^0 15^5 Mid-cauline 18^10 leaf length Upper densely glandular and densely glandular usually densely glandular, internode moderately villous can be glabrous pubescence Lateral 4-7 flowers/cyme, 6-11 flowers 3-5 flowers/cyme cymes ^MIK- i i. l.lkl.ll hl.llljl mm mm mm Calyx 6-7 5-7 4-6 long, long, long, mm mm mm dimensions lobes 2-A (—6) lobes 1-2 lobes 3.5-5 Calyx color tube ribs green and/or tube white ventrally, tube ribs green and purple, membrane membrane purple, purple dorsally; connecting ribs connecting ribs white lobes green white, sometimes is is or purple; lobes green purple; lobes purple, green distally mm mm mm Calyx awn 0.2-0.3 0.5-1 0.6-0.9 length Corolla corolla slightly tube and lobes pink-to- corolla satin; tube white. color translucent; tube hot pink, never dotted or pink, hot pink, salmon, heavily coral pink streaked lavender, tints of blue; streaked; lobes dotted lobes sometimes dotted and streaked coral pink Corolla moderately to heavily glabrous lightly to moderately pubescence glandular glandular mm mm mm 14-17 15-18 25-45 Corolla tube length (re: variability of style length/anther position) > Schneider and Bregar: Ipomopns ramosa sp. nov. { , mm mm mm 6-8 Corolla acute, 8 long obtuse, long acute, 11 long lobes mm mm mm Style length 2.5-4 1.3-2 16-35 Stigma below the anthers below the anthers below anthers to slightly position at (reverse herkogamous) (reverse herkogamous) exserted (approach anthesis herkogamous to reverse herkogamous) Stamens unequally inserted 5-10 unequally inserted 4-6.5 stamens unequally inserted mm mm mm above base of tube, above base of tube, within 6 of orifice of 0-5 anthers well included anthers included tube, anthers mm i-wi U-»l 1" 2 mm mm mm Filaments 0.1-1 long, straight 0.4-0.6 long 1-1.5 (often kinked) Pollen yellow light yellow to bluish white to yellow, rarely blue Seeds 2-3 per locule, the seed 2-4 per locule, the seed unknown casing becomes casing becomes mucilaginous when wet mucilaginous when wet '"Details about tenuituba are from a variety of sources (personal measurements of plants in the /. & may Abajo Mountains; Welsh 2008; Cronquist 1984; Wilken 1993, Grant Wilken 1988) and et al. not be exact for all populations of this highly variable species over its entire range. ACKNOWLEDGEMENTS We are grateful to Dieter Wilken and Mark Porter for their comments on the penultimate J, We manuscript. thank Bob Sivinski and Phil Tonne for sharing Ipomopsis sancti-spiritus information and photographs, Charlie McDonald for making the 1988 Wilken and Fletcher description of Ipomopsis sancti-spiritus available to us, Bob Powell for helpful comments on the initial manuscript, and Kim Feemen for making us aware of the second population of Ipomopsis ramosa in Wildcat We Many Guy Canyon. thanks to Betty for assisting with her sharp eyes. are especially grateful to Nesom for expeditiously providing suggestions, editorial assistance, and the Latin translation. Photographs are by the authors; also see Schneider (201 for more photographs. 1) LITERATURE CITED A A Cronquist, 1984. Pp. 86-155, in Cronquist, A.H. Holmgren, N.H. Holmgren, J.L. Reveal, and P.K Holmgren (eds.). Vascular Plants of the Intermountain West, Vol. 4. New York Botanical Garden Press, Bronx. CU Museum-COLO. Museum Specimen Database of Colorado Vascular Plants. Univ. of Colorado of Natural History, Boulder. <http://cumuseum.colorado.edu/Research/Botany/ php> Databases/search. Grant, V. and D.H. Wilken. 1988. Racial variation in Ipomopsis tenuituba (Polemoniaceae). Bot. Gaz. 149: 443^149. New Mexico Rare Plant Technical Council (NMRPTC). 2011. New Mexico Rare Plants Website. Albuquerque. <http ://nmrareplants. unm edu> . Rocky Mountain Herbarium. 2011. Rocky Mountain Herbarium Specimen Database. Univ. of Wyoming, Laramie, <http://www.rmh.uwyo.edu/data/search.php> & Schneider, A. 2011. Southwest Colorado Wildflowers, Ferns, Trees. For identification and <www com appreciation. swcoloradowildflowers. . SEINET. 2011. Southwest Environmental Information Network. Managed at Arizona State Univ., Tempe, <http://swbiodiversity.org/seinet/iridex.php> W.A Weber, and R.C. Wittmann. 2001. Colorado Flora: Western Slope (ed. 3). Univ. Press of Colorado, Boulder. Schneder ^nd Brtgar: Ipomopsisi Welsh, SL. N.D. Atwood, S. Goodrich, and L.C Higgins. 2003. A Utah Flora (ed. 4). Print Services, Bngham Young Univ., Provo, Utah m Wilken, D.H. 1993. Polemoniaeeae. Pp. 836-339, J.C. Hickman (ed.) Jepson Manual: Higher Plants of California. Univ. California Press, Berkeley Wilken, D H. and R Fletcher. 1938. Ipomopsis sancti-spiriOis (Polemoniaeeae), a new species from northernNew Mexico Brittonia 40: 48-51. Wilken, D.H. and J.M. Porter. 2005. Vascular Plants of Arizona' Polemoniaeeae. Canotia 1: 1-37. <http://nhc.asu.edu/vpherbanum/canotia/CANariA-29Nov05-voll.pdf> Figure 1 High above the Roaring Fork Creek, the road climbs 1500 feet along the canyon vails which are the type locale for Ipomopsis ramosa. The plant is found intermittently along the first 5 miles ofthe road as climbs eastward from Colorado Highway 145. it 5chne)der and Bregar: Spomopsisramosa,sp. nov. Figure 2. Typical rocky, steep, mountainsidehabitat andtypical spacing for Ipomopsis ramosainthe Roaring Fork Canyon, thetypelocality. mm mm Figure 4. Corollas ate slightlytranslucent;tubes ara 14-17 long, 1 wide attheirbase inside mm the calyx, and 2 wide at the orifice where the bbes flair; tubes are heavilystripedcoral pink; mm lobes ara 8 long, recurved, coral-pinkdottedon theventral surface, andmore heavilycoral-pink streakedonthe dorsal surface. Fromadistance,the corollasappearsalmon or orange,but aclose-up examination showsthemtobe coralpinkandwhite. The center area ofthe tube mayappearyellow but thatis due to the translucencythatallows the yellowanther color to showthrough. Calyces ara stronglymulti-coloredand, consistentwiththe rest oftheplant stronglyglandular.